The Theropod Ancestry of Birds: New Evidence from the Late Cretaceous of Madagascar C. A. Forster, S. D. Sampson, L. M. Chiappe, and D. W. Krause

Character matrix for phylogenetic analysis of Rahona ostromi.

                    5     10     15     20     25     30     35     40     45
                   50     55     60     65     70     75     80     85     90
                   95    100    105    110    113

Allosaurus     00000  00000  00000  00000  00000  00000  00000  00000  00000
                00000  00000  00000  00000  00000  00000  00000  00000  00000
                00000  00000  00000  00000  000

Compsognathus  0100?  00000  10???  ?0?11  01??0  ?00?0  ??00?  00??0  ?0000
                ?0000  ???0?  ?0000  00??0  0?00?  000??  1?100  10?00  0??0?
                ?????  ?????  ?00??  00000  1?0

Archaeopteryx  02111  00111  10110  01?11  01000  100?1  00101  1?101  10010
                11010  01110  00010  01112  10111  11001  11111  12110  11000
                111??  0111?  ?0111  00100  100

Rahona         ?????  ?????  ?????  ?????  ?????  ??111  11?01  10??1  10011
                ????0  ?????  ??01?  ????2  ??11?  11101  11111  12110  11000
                21200  01112  11010  0?101  100

Oviraptoridae   1???1  10101  10110  00001  01001  00101  10000  00000  10000
                10000  01100  00000  00001  00001  00000  00000  11000  00000
                11000  00000  00000  00000  100

Dromaeosauridae 01001  10001  00110  11011  01000  00101  00001  10001  00000
                10010  01010  00000  01111  00011  00000  10110  11000  01000
                11100  00000  00010  00001  000

Ornithomimidae  02011  11010  10110  10111  01001  00000  00000  00010  00000
                10000  00000  00000  00011  00000  00000  00010  11000  00000
                11000  00000  00001  00010  000

Troodontidae    00011  01111  001?0  ?0111  11001  ?0100  10001  10011  00000
                10000  00?10  00010  01011  10011  0000?  ?0011  11000  01000
                11200  00001  00110  0?011  100

Tyrannosauridae 00001  11101  00000  00000  10010  00001  00000  00000  00000
                00000  00000  00000  00020  00000  00000  00100  10000  00000
                00000  00000  00000  00010  000

Alvarezsauridae ?1111  01?10  11??1  11011  1?001  ?0110  11101  10110  11000
                10000  00211  00001  0??20  11101  0??1?  02201  2??00  00211
                21000  00111  11110  00?10  100

Enantiornithes  01111  01?10  111??  1???1  ????1  ?1110  11211  ?111?  10111
                21120  12211  11111  11112  00111  00111  0?201  2?101  11100
                21?11  1011?  11211  01100  100

Ornithurae      11111  11110  11021  1?011  11111  11110  11211  ?111?  10101
                21121  12211  11111  11112  01111  00211  02201  2?001  11211
                21011  10112  11211  11100  111

Unenlagia      ?????  ?????  ?????  ?????  ?????  ???01  11???  ?0???  ??01?
                ????0  ????0  0????  ?????  ?????  11001  11111  11100  11000
                11000  01???  ?????  ?????  ???

Patagopteryx   ?????  ?1?10  ????1  1???1  ??11?  ?1110  11???  ?111?  11111
                21121  1?11?  ??011  ??0??  0111?  00010  02201  20001  10211
                210?1  10011  11211  11000  111

Iberomesornis  ?????  ?????  ?????  ?????  ?????  ?01?0  0011?  ???1?  101??
                21?20  12???  00?1?  ?????  ???1?  0????  ??1??  2?10?  1????
                ?1???  ??1??  ????0  0?100  1?0

Character List for phylogenetic analysis of Rahona ostromi

1. teeth in premaxilla: present (0); absent (1).
2. denticles on teeth: present on anterior and posterior carinae (0); present on posterior carina, but absent or severely restricted on anterior carina (1); absent (2). (adapted from Martin et al., 1980)
3. shape of teeth: laterally compressed and recurved (0); slightly compressed, nearly conical (1). (Ostrom, 1976)
4. constriction between crown to root: absent (0); present (1). (Martin et al., 1980)

5. jugal contribution to antorbital fenestra: absent, eliminated by lacrimal-maxilla contact (0), present (1). (Holtz, 1994)
6. maxilla contribution to naris: absent, excluded by a nasal-premaxilla contact (0); present (1). (Martin et al., 1980; Currie, 1995)
7. shape of frontals: anteriorly broad (0); triangular, narrow rostrally (1). (Currie, 1987b; Russell and Dong, 1993)
8. extreme reduction or loss of prefrontal: absent (0); present (1). (Gauthier, 1986)
9. frontal length relative to parietal: smaller or subequal (0); nearly two times as long (1).
10. narrow sagittal crest on parietals: absent (0); present (1). (Currie and Zhao, 1993)
11. jugal bar shape: robust (0); thin and straight (1). (Ostrom, 1976)
12. jugular postorbital process: present (0); absent (1). (Chiappe)
13. subsidiary foramen between palatine and pterygoid: absent (0); present (1). (Gauthier, 1986)
14. palatine-ectopterygoid contact: no contact, ectopterygoid present (0); contact present, ectopterygoid present (1); no contact, ectopterygoid absent (2). (adapted from Currie, 1995)
15. medial otic process of quadrate that articulates with prootic: absent (0); present (1). (Chiappe, 1995)
16. distal quadrate articulation set well anterior to proximal quadrate cotyle: absent (0); present (1). (Ostrom, 1976)
17. shape of paroccipital process: straight and vertically oriented (0); distal end with distinct twist as to face caudodorsally (1). (Colbert and Russell, 1969; Currie, 1995)
18. bulbous parasphenoid capsule (cultriform process): absent (0); present (1). (Osmolska et al., 1972)
19. accessory lacrimal fenestration: present (0); absent (1).
20. supraoccipital crest: absent or weak (0); well pronounced (1).
21. fusion between parietals and laterosphenoid in adults: present (0); absent (1).
22. accesory fenestra between premaxilla and maxilla: present (0); absent (1).
23. quadratojugal cotyla in the lateral face of the mandibular process of the quadrate: absent (0); present (1).
24. pneumatic quadrate: absent (0); present (1).

25. unfused interdental plates: present (0); absent (1). (Currie, 1987; Holtz, 1994)
26. separate coronoid bone: present (0); absent (1). (Elzanowski and Wellnhofer, 1996)

27. heterocoelus cervical vertebrae: absent (0); present (1). (Sanz et al., 1995)
28. ventral processes (hypapophyses) on cervicodorsal vertebrae: absent or very weakly developed (0), present, well developed (1). (Gauthier, 1986)
29. ratio of height of neural canal in dorsal vertebrae to height of cranial articular face: less than .40 (0); more than .40 (1). (Sanz et al., 1995)
30. pneumatic foramina on dorsal vertebrae: absent (0); present (1). (Ostrom, 1976)
31. number of sacral vertebrae: 5 or less (0), 6 or more (1). (adapted from Ostrom, 1976)
32. fusion of all sacral vertebrae in adult: absent or partially fused (0); present, all completely fused (1).
33. number of caudal vertebrae: 30 or more (0), 20-25 (1), or less than 15 (2).
34. pygostyle: absent (0); present (1) (Sanz and Buscalioni, 1992)
35. transition point on caudal vertebrae: distal, behind caudal 10 (0); proximal, no further back than caudal 10 (1) (Gauthier, 1986; Gatesy, 1996)
36. middle and distal chevron shape: laterally compresses, expanded dorsoventrally (0); dorsoventrally flattened into a thin horizontal plate behind transition point (1). (adapted from Ostrom, 1976; Gauthier, 1986)
37. hyposphene-hypantrum articulations on dorsal vertebrae: present (0); absent (1).
38. pneumatic foramen on sacral vertebrae: present(0); absent (1).
39. cervical neural spines: dorsoventrally tall, axially short (0); dorsoventrally short, axially elongate (1).
40. length of middle and posterior caudal vertebrae: subequal to length of proximal caudals (0); elongate, at least 130% the length of the anterior caudals (1).
41. prezygopophyses on middle and posterior caudal vertebrae: elongate, extend at least half the way across preceeding vertebral centrum (0); reduced or absent, extend over 25% or less the length of preceeding vertebral centrum (1).
42. well-developed procoely in anterior caudal vertebrae: absent (0); present (1).

43. scapula with tapered distal end: absent (0); present (1). (Ostrom, 1976)
44. anterior projection of acromion process: absent (0); present, extends well anterior to glenoid fossa (1).
45. scapulocoracoid articulation: immobile, sutural (0); mobile, cartilagenous (1) (adapted from Sanz et al., 1995)
46. acroracoid of coracoid: absent, biceps tubercle small (0); absent, biceps tubercle large (1); present, extends above scapulocoracoid articulation, biceps tubercle large (1). (Perle et al., 1993; Walker, 1972; Ostrom, 1976)
47. body of coracoid forming sharp angle with the body of the scapula: absent (0); present (1) (Ostrom, 1976; Tarsitano and Hecht, 1980)
48. long axis of coracoid portion of glenoid relative to long axis of scapular portion of glenoid: alligned (0); form a 90 degree angle (1). (Chiappe)
49. coracoid shape: short, rounded sternal border (0); elongate, subrectangular profile, rounded sternal border (1); elongate, strut-like (2). (adapted from Ostrom, 1976; Gauthier, 1986)
50. dorsoventrally curved scapular shaft: absent (0); present (1).
51. supracoracoid nerve foramen (incision) located in the medial margin of coracoid: absent (0); present (1). (Sanz et al., 1995)
52. shape of fused clavicles: absent (0); present, interclavicular angle greater than 90 degrees (1); present, interclavicular angle less than 90 degrees (2). (Ostrom, 1976; Chiappe, 1991)
53. ossified sternum: paired (0); fused or partially fused into a single structure (1); fused with a midline keel (2).

54. distal condyles of humerus located mainly on the distal (0), or cranial (1), aspect. (Perle et al., 1993)
55. ventral tubercle of humerus projects distinctly caudally, separated from the humeral head by a deep capital incision: absent (0); present (1) (Sanz et al., 1995)
56. well-developed transverse ligamental groove on the humerus: absent (0); present (1) (Sanz et al., 1995)
57. humerus length relative to ulna: longer (0); subequal to or shorter (1). (Perle et al., 1993)
58. dorsal condyle of ulna developed as a semilunar ridge: absent (0); present (1) (Sanz et al., 1995)
59. ratio of diameter of shaft of radius to ulna: more than .70 (0); thinner than ulna, less than .70 (1). (adapted from Tarsitano and Hecht, 1980; Perle et al, 1993; Chiappe and Calvo, 1994)
60. carpometacarpus: absent (0); present (1). (Sanz et al., 1995)
61. extensor process on metacarpus: absent (0); present (1). (Sanz et al., 1995)
62. metacarpal I greater than (0), or less than (1) one-third the length of MC II. (Tarsitano and Hecht, 1980; Gauthier, 1986)
63. metacarpal III: straight (0); bowed laterally (1). (adapted from Tarsitano and Hecht, 1980; Gauthier, 1986)
64. combined lengths III-1 and III-2 relative to III-3 on manus: longer (0); subequal (1); absent (2). (Gauthier, 1986)
65. forelimb elongatation relative to presacral length: less than one half presacral length (0); 60-90% presacral length (1); more than 100% presacral length (2). (adapted from Ostrom, 1976; Gauthier, 1986)
66. manus length relative to ulna length: manus shorter than or subequal to ulna (0); manus longer than ulna by at least 20% (1).
67. proximal and distal ends of humerus: twisted, do not lie in the same plane (0); expanded into the same plane (1).
68. ulnar distal condyle: transversely compressed and craniocaudally extended proximally in the same plane as the humero-ulnar flexion-extension movement (0); subtriangular in shape in distal view, with a dorsomedial condyle, and twisted more than 54 degrees with respect to the proximal end (1).
69. shape of ulnar posterior margin: sigmoid (0); uniformly convex (1).
70. distal radial carpal: proximodistally flattened (0); semilunate (1).

71. length of preacetabular process of ilium relative to length of postabetabular process: subequal (0); twice as long (1). (adapted from Tarsitano and Hecht, 1980; Currie and Russell, 1988)
72. postacetabular process depth: relatively deep, more than 50% depth a acetabulum (0); shallow, less than 50% depth at acetabulum, drawn back into a low, pointed process (1). (adapted from Ostrom, 1976; Novas, 1997)
73. ratio of lengths of acetabulum to ilium: .15 or more (0); .11 to .13 (1); .10 or less (2). (adapted from Sanz et al., 1995)
74. antitrochanter on ilium: absent or very small (0); present, prominent (1). (adapted from Ostrom, 1976; Perle et al., 1993)
75. tubercle on dorsal margin of ilium above caudal acetabulum: absent (0); present (1).
76. craniocaudal width of pubic peduncle on ilium relative to width of acetabulum: narrow, less than acetabular width (0); wide, exceeds acetabulum in width (1).
77. pubic foot: projects cranially and caudally (0); projects caudally only (1); absent (2). (adapted from Ostrom, 1976; Gauthier, 1986; Sanz et al., 1995)
78. angle of pubic shaft relative to the long axis of sacral vertebrae: projects cranially (0); subvertical (1); projects caudally (opisthopubic) (2).
79. ratio of length of ischium to length of pubis: more than .66 (0), less than .66 (1). (adapted from Ostrom, 1976; Gauthier, 1986)
80. contact between distal ischia: present (0); absent (1). (Tarsitano and Hecht, 1980; Sanz et al., 1995)
81. obturator process on ischium: present, square (0); present, peaked and broadly triangular (1); rudimentary or absent (2).
82. placement of obturator process: proximal (0); mid-shaft (1); distal (2).
83. "postacetabular" process on proximal ischium behind iliac contact: absent (0); present, nearly contacts postacetabular process of ilium (1). (Ostrom, 1976)
84. process on caudal border of midshaft of ischium: absent (0); present (1). (Ostrom, 1976)
85. fusion of pelvic elements: absent (0); present (1). (Sanz et al., 1995)
86. shape of ischial shaft: rod-like in part with a circular or subcircular cross section (0); mediolaterally compressed and plate-like along entire length (1).
87. postacetabular blade on ilium: brevis shelf caudolaterally oriented, medial flange ventrally curved (0); postacetabular blade vertical, medial flange strongly reduced and perpendicular to iliac blade (1). (Novas, 1997)
88. pubic apron: transversely broad, pubes fused for distal 2/3rds of their length (0); transversely narrowed, pubes fused only along the distal 1/3rd of their length (1).
89. contact between distal pubes: present (0); absent (1).
90. iliac fossa for m. cupedicus: present (0); absent (1).

91. configuration of lesser trochanters: large and flange-like, separated from femoral shaft, head, and greater trochanter by a deep cleft (0); nearly confluent with the greater trochanter, and separated from the proximal head of femur by only a small cleft or groove (1); joined to greater trochanter to form an undivided trochanteric crest, proximal articular surface confluent with that of femoral head (2). (adapted from Ostrom, 1976; Tarsitano and Hecht, 1980; Gauthier, 1986; Currie and Russell, 1988; Perle et al., 1993)
92. shape of fourth trochanter on femur: present, large (0); reduced to a low ridge or absent (1). (adapted from Tarsitano and Hecht, 1980; Gauthier, 1986; Chiappe, 1996)
93. "posterior trochanter" on femur: absent (0); present, cranially placed (1); present, centered on the trochanteric crest (2). (adapted from Ostrom, 1986; Sanz et al., 1995)
94. fossa for capital ligament: absent (0); present (1).
95. popliteal fossa of femur bounded distally by transverse bridge: absent (0); present (1). (Perle et al., 1993)
96. tibiofibular crest on the lateral condyle of distal femur: absent (0); present (1). (Sanz et al., 1995)
97. femoral neck: present, constriction developed that separates trochanteric region from femoral head (0); absent (1) (adapted from Russell, 1969)
98. proximal tibia: craniocaudal length twice that of mediolateral width (0); length and width of proximal tibia subequal (1).
99. midshaft diameter of fibula relative to tibia: approximately one-fifth or more that of tibia (0); one-fifth or less that of tibia (1).
100. orientation of iliofibularis tubercle on fibula: craniolateral (0); lateral (1); caudolateral or caudal (2). (Forster et al., 1996)
101. deep fossa on medial side of proximal fibular head: present (0); absent (1).
102. fibular articulation with the calcaneum: present (0); absent, fibula does not reach tarsus (1)
103. fusion of proximal tarsals to crus: absent (0); present, partially fused (1); present, completely fused (2). (adapted from Ostrom, 1976; Forster et al., 1996)
104. fusion of astragalus to calcaneum in adults: absent (0); present, partial to complete (1). (Currie and Peng, 1993)
105. fusion of distal tarsals to metatarsus: absent (0); present (1). (Ostrom, 1976)
106. fusion of metatarsus: absent or partially fused (0); completely fused (1). (Ostrom, 1976)
107. metatarsal V: present (0); absent (1). (Ostrom, 1976)
108. position of pes digit I: parallel to other digits (0); reversed to oppose other digits (1). (Ostrom, 1976)
109. relative contributions of metatarsals II, III, and IV to ankle joint: all contribute approximately equally (0); partially or completely arctometatarsalian, MT III is nearly or completely eliminated from joint (1). (Currie and Peng, 1993)
110. relative size of pes digit II: phalanges and ungual subequal in size and robustness to digits III and IV (0); developed into a robust, hyperextendable slashing digit with an enlarged sickle-like ungual (1). (Gauthier, 1986)
111. ratio of length of tibia to length of femur: tibia no more than 15% longer than femur (0); tibia elongate, at least 25% longer than femur (1).
112. extensor canal on tibiotarsus: absent (0); present (1).
113. distal vascular forament on metatarsus (tarsometatarsus): absent (0); present (1).

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